Order DERMAPTERA

Dermaptera

Museums

Museum list

Regional Maps

Order DERMAPTERA

Earwigs

Compiler and date details

2006 - updated ABRS, data provided by Fabian Haas

1998 - Gerasimos Cassis, Australian Museum, Sydney, New South Wales, Australia

Introduction

The Dermaptera are a moderate-sized order of exopterygote neopteran insects. This uniform group of terrestrial insects, commonly known as earwigs, are distinctive because of their forceps-like cerci. They are often cryptic, living under bark and stones, amongst litter and in refuse and composts.

Earwigs are elongate, flattened or cylindrical, winged or apterous. The head is prognathous, with mandibulate mouthparts, and usually with large eyes. The thorax is flexible and has free segments. The pro- and mesothorax are subequal in size and the metathorax is enlarged; in winged species the metanotum has a tegminal lock process. The forewings are short and lack veins. The hind wings are large, semicircular, mostly composed of a vannal fold, and are folded fan-like; many species, particularly of the Anisolabididae, are apterous. The legs are cursorial and the tarsi are three-segmented. The abdomen is elongate and flexible, with 10 visible segments in the male and eight in the female. The cerci of the Forficulina are forcipate, usually heavily sclerotised, unsegmented, and often sexually dimorphic, with male forceps usually more arcuate. The male genitalia, integral to the classification of earwigs, consist of a subcylindrical intromittent organ, which is forked apically, with either a single median genital lobe, or two lateral lobes. The female ovipositor is greatly reduced in all Forficulina, except for the Pygidicranidae, and some Apachyidae and Labiduridae-Allostethinae, which have rod-like valves. The larvae resemble adults, with the wings absent and the forceps simplified (Giles 1963; Brown 1982; Rentz & Kevan 1991).

Giles (1963) reviewed earlier views on the affinities of the Dermaptera and concluded that they are most closely allied to the Grylloblattodea, and that both are considered to be basal in the orthopteroid complex of orders. Tillyard (1931) and Crampton (1933) were in agreement that the earwigs are descended from the stem-group, Protelytroptera. This hypothesis was supported by Kukalová-Peck & Peck (1993), who also recognised the Zoraptera as the sister-group of the Dermaptera + Protelytroptera. Additionally, they supported Kukalová-Peck's (1991) review of the classification of the exopterygote Neoptera, and they concluded that the Dermaptera belong to the blattoid assemblage, the Blattoneoptera, and not to the orthopteroid orders. Kristensen (1991) indicated that the Dermaptera may have affinities with the Plecoptera and Embioptera, primarily on the basis of genitalic characters, and with the Phasmatodea, due to similarities in sperm specialisations (also Jamieson 1987). Whiting et al. (1997) proposed a sister-group relationship between the Plecoptera and Dermaptera on the basis of molecular and morphological data. Rentz & Kevan (1991) implied that the relationships of Dermaptera are obscure by reporting the presence of small and holocentric chromosomes, in comparison to large, monocentric chromosomes in other blattoid and orthopteroid orders (also Webb 1970).

CLASSIFICATION

Higher classification of the Dermaptera

The higher classification of the Dermaptera remains somewhat unresolved. Conflicting modern classifications have been proposed by Popham (1965–1990), Steinmann (1986–1993), Sakai (1970–1997) and Haas (1995) (Table 1). The Dermaptera, as currently accepted, comprises three extant suborders: Hemimerina, Arixeniina and Forficulina. The extinct suborder Archidermaptera, regarded as the sister-group of the extant suborders (Willmann 1990), is known from the Jurassic (Kukalová-Peck 1991) and contains ten species, all of which have adults with segmented cerci.

Table 1. Higher classification of the Dermaptera by major authors. Family-group names are given as in the original works. Incorrect subsequent spellings are not corrected in the table. Comments on the classifications of each author, and comparisons, are given in the following text.

Verhoeff (1902)
EUDERMAPTERA
DIANDRA
Gonolabidae
Anisolabidae
Labiduridae
Pygidicranidae
Diplatyidae
MONANDRIA
Karschiellidae
Cheliduridae
Forficulidae
PARADERMAPTERA
Apachyidae

Zacher (1910–1915)
PROTODERMAPTERA
PYGIDICRANIALES
Diplatyidae
Pygidicranidae
Echinosomatidae
Karschiellidae
LABIDURIALES
Allostethetidae
Anisolabidae
Brachylabidae
Labiduridae
Esphalmenidae
PARADERMAPTERA
Apachyidae
EUDERMAPTERA
Labiidae
Cheliduridae
Forficulidae

Burr (1909–1916)
HEMIMERINA
ARIXENIINA
FORFICULINA
PROTODERMAPTERA
Pygidicranidae
Labiduridae
(Apachyinae)
Psalidae
EUDERMAPTERA
Labiidae
Chelisochidae
Forficulidae

Popham (1965–1990)
HEMIMERINA
FORFICULINA
PYGIDICRANIDEA
Pygidicranidae
FORFICULIDEA
LABIOIDEA
Carcinophoridae
Arixenidae
Labiidae
FORFICULOIDEA
Labiduridae
(Apachyinae)
Chelisochidae
Forficulidae

Sakai (1970–1997)
HEMIMERINA
ARIXENIINA
FORFICULINA
PYGIDICRANOIDEA
Pygidicranidae
Diplatyidae
ANISOLABIDOIDEA
Anisolabididae
Labiduridae
APACHYOIDEA
Apachyidae
FORFICULOIDEA
Spongiphoridae
Forficulidae
Chelisochidae

Steinmann (1973–1993)
?HEMIMERINA
?ARIXENIINA
CATADERMAPTERA
PROTODERMAPTERA
PYGIDICRANOIDEA
Pygidicranidae
MESODERMAPTERA
Carcinophoridae
Labiduridae
PARADERMAPTERA
APACHYOIDEA
Apachyidae
EUDERMAPTERA
FORFICULOIDEA
Labiidae
Chelisochidae
Forficulidae

Haas (1995)
HEMIMERINA
ARIXENIINA
FORFICULINA
Karschiellidae
Diplatyidae
Pygidicranidae
Apachyidae
Labiduridae
Anisolabididae
Spongiphoridae
Chelisochidae
Forficulidae

In all but early works of Popham, the epizoic Hemimerina are regarded universally as a dermapteran suborder. Burr (1911) erected a new suborder, the Arixeniina, for the bat commensals of the Old World Tropics, and this has been followed by most authors. Popham (1961) disputed this arrangement, subsequently placing them within the Forficulina-Labioidea, as the sister-group to the Labiidae, on the basis of gut and male genitalic characters (Popham 1985). Giles (1974) argued for independent derivations of the Hemimerina and Arixeniina from forficuline stock, and subordinal status for both taxa. Steinmann (1973–1993) made almost no mention of the Hemimerina and Arixeniina, although in his later works he implicitly removed them from the Dermaptera.

All major authors accept a single suborder for the free-living earwigs, the Forficulina. The infraordinal, sectional and superfamily classifications of these authors differ in hierarchical arrangement, representative families and sister-group relationships.

Verhoeff (1902) gave the first phylogenetic classification of the earwigs, recognising two suborders: the Paradermaptera for the Apachyidae, and the Eudermaptera for the remaining families. He divided the Eudermaptera into eight families: the Diplatyidae, Pygidicranidae, Labiduridae, Anisolabidae (sic), Gonolabidae (sic), Cheliduridae, Karschiellidae and Forficulidae.

Zacher (1910b) accepted the Paradermaptera as an infraordinal category for the Apachyidae, although he placed them as the sister-group to his Esphelmatidae, within the suborder Protodermaptera. Later, he was less certain about this arrangement (Zacher 1911). He restricted the Eudermaptera to the Labiida (sic), Cheliduridae and Forficulidae. He also regarded the Apachyidae as a distinct group, the Paradermaptera. The remaining families he grouped in his suborder, Protodermaptera, which contained the Pygidicraniidae, Labiduridae and related families.

Burr's (1909–1916) classification of the Forficulinae recognised three superfamilies: Protodermaptera (Pygidicranidae, Labiduridae), Paradermaptera (Apachyidae) and Eudermaptera (Labiidae, Chelisochidae and Forficulidae), with the subfamilial ranking of the Diplatyinae, Karschiellinae and Brachylabinae (sic) (Pygidicranidae), Esphalmeninae and Allostethinae (Labiduridae), taxa which were accorded family status by either Verhoeff or Zacher. Within his series of works, Burr arranged and rearranged his classification of the Dermaptera, particularly his ranking and placement of Apachyidae and ranking of Chelisochidae.

Popham's (1965–1990) classification is the most eccentric. Initially, he dispensed with the Protodermaptera, Paradermaptera and Eudermaptera, and recognised four superfamilies and nine families: Labioidea (Carcinophoridae, Arixenidae, Labiidae), Forficuloidea (Labiduridae, Chelisochidae, Forficulidae), Pygidicranoidea (Diplatyidae, Pygidicranidae) and Karschielloidea (Karschiellidae) (Popham 1965a, 1965b, 1965c). Subsequently, he revised his classification (Popham 1985–1990), recognising two suborders, Hamimina and Forficulina, with the latter divided into two infraorders, the Pygidicranidea and Forficulidea, the former restricted to the Pygidicranidae (including Karschiellinae and Diplatyinae), and the latter containing all other families in two superfamilies, the Labioidea (sic) (Carcinophoridae, Arixenidae, Labiidae) and Forficuloidea (Labiduridae, Chelisochidae, Forficulidae). Popham's classification was originally based on the male genitalia, particularly on the presence of the basal vesicle of the virga, but later included a number of external characters (cf. Popham 1965c and 1985). Popham (1997) commented on the various classifications without providing any synthesis contrary to his previous classification.

Sakai's (1970–1997) classification recognises four superfamilies and supports the Mesodermaptera (as Anisolabididoidea) as a distinct taxon. Additionally, he has consistently regarded the Diplatyidae as distinct from the remainder of the Pygidicranoidea. The justification for his classification is wanting, although it can be seen as syncretic, and is most like Steinmann's classification.

Steinmann's (1973–1993) classification is essentially a further development of Burr's classification, although he introduced a new suborder, the Catadermaptera, and new infraordinal and sectional categories—the Mesodermaptera, for the Carcinophoridae and Labiduridae. His comprehensive work accounts for all described earwig species and is more synthetic, with the inclusion of external morphological characters and male genitalia. His critique of previous schema questions the ranking of genus and family-groups of other workers, although the rationale for his arrangement is inadequate.

Like Popham, Haas (1995) provided a phylogenetic classification, which is based in part on a re-assessment of the characters of others, including Giles (1963). Haas concluded that the monophyly of certain suprafamilial taxa of Popham and Steinmann is not supported, particularly the Labioidea and Mesodermaptera. Furthermore, he concluded that the Karschiellidae warrant family rank, excluded diplatyines from the Pygidicranidae sensu stricto, and supported the family ranking of the Apachyidae. Haas also discussed the sister-group relationships of dermapteran families, and rejected many of the sister-groups proposed by Popham (1985).

Implicit in Haas' (1995) work is the rejection of any suprafamilial classification of the Forficulina. Due to the lack of comprehensive analysis of characters and the restricted taxonomic sample of previous works, I have adopted Haas' cautious schema, even though it departs from the arrangement in Insects of Australia (Rentz & Kevan 1991), which follows Sakai (1970–1997).

Familial classification of the Dermaptera
In Cassis (1998) the familial classification, follows the arrangement of Haas (1995, see Table 1), recognising nine families, two of which have not been recorded from Australia. The Karschiellidae are endemic to the Afrotropical Region and are represented by two genera: Karschiella Verhoeff and Bormansia Burr and twelve species (Steinmann 1986; Sakai 1997). The Diplatyidae are a distinctive taxon, with a circumtropical distribution and represented by five genera and 161 species (Sakai 1997). Steinmann (1986, 1989a), amongst others, has considered these taxa to be subfamilies of the Pygidicranidae. Sakai (1987) and Haas (1995) argue strongly for their separate status—Haas disputing the monophyly of the Diplatyidae—on the basis of differences of wing character states between Haplodiplatys Hincks and the remainder of the Forficulina.

The Pygidicranidae are uniformly regarded as the most plesiomorphic family of the Forficulina. The family includes 20 genera (Steinmann 1989a) and 203 species (Sakai 1997; note that Sakai does not include numbers of genera, which may account for discrepancies in taxon richness between his and Steinmann's classifications), with a largely circumtropical distribution (Popham 1963; Steinmann 1989a). Haas (1995) has indicated doubt about the monophyly of the Pygidicranidae sensu stricto. Alternative subfamilial classifications (cf. Steinmann 1986, 1989a; Sakai 1997) exist, and are discussed in the chapter on the Pygidicranidae.

The position and rank of the Apachyidae remains contentious in the family-group classification of the earwigs. Burr (1915a) and Popham (1985) regarded this taxon as a subfamily of the Labiduridae in contrast to all other authors who support their familial status. Apachyidae monophyly is firmly established by the presence of a squamopygidium. The family is composed of two genera, Apachyus Audinet-Serville and Dendroiketes Burr, and 15 species found under the bark of trees in the Eastern Hemisphere—most are restricted to the tropics.

The composition of the Labiduridae in this work, follows that of Steinmann (1989a, 1989b), Haas (1995), Sakai (1997) and others, and is restricted to the subfamilies Allostethinae, Nalinae and Labidurinae. The family is composed of seven genera and 71 species (species numbers are based on Sakai (1997)). The majority of species are known from the tropics of the Oriental Region. The complex synonymy of the tramp species, Labidura riparia (see Sakai 1987) remains contentious (discrepancies between Sakai's and Steinmann's classifications derive from differences in this synonymy), and the endemic Labidura truncata Kirby, is here considered a distinct species. Knowledge of the biology of L. riparia and of the economic pest, Nala lividipes (Dufour), constitute much of what we know about labidurid biology (Simpson 1989).

The Anisolabididae are a large, complex family of earwigs, many of which are apterous. The family is composed of nine subfamilies, 33 genera and 349 species (Sakai 1997) and has a cosmopolitan distribution. Sakai (1997), however, recognised only eight subfamilies, not the Gonolabinae Popham & Brindle, 1966. The majority of species belong to the Anisolabidinae, which contains the speciose genera, Anisolabis Fieber, Carcinophora Scudder, Gonolabis Burr and Euborellia Burr, and the broadly distributed tramp species, Euborellia annulipes (Lucas) and Anisolabis maritima (Bonelli). Haas (1995) has questioned the monophyly of the Anisolabididae, although he does not give any clarification as to which groups are problematic.

The Spongiphoridae are the most diverse earwig family, comprising 13 subfamilies, 39 genera and 512 species (Steinmann 1990; Sakai 1997) and are cosmopolitan in distribution. The adults are usually small and winged. Steinmann (1990) concluded that the family is inadequately diagnosed by the presence of the simple second tarsus. He suggested that the suprageneric classification of the family is 'extremely problematic'. There are a number of speciose genera, including Chaetospania Karsch, Spongovostox Burr, Marava Burr and Paralabella Steinmann.

The Chelisochidae are represented by three subfamilies, 16 genera and 96 species (Steinmann 1993; Sakai 1997—recognised only two subfamilies, Chilisochellinae and Chelisochinae, not Genitalatinae Steinmann, 1987). They are mostly restricted to the Eastern Hemisphere, particularly the tropical regions, except for the tramp species, Chelisoches morio (Fabricius) which has a circumtropical distribution.

The Forficulidae are a distinctive cosmopolitan family, comprising eight subfamilies, 65 genera and 490 species. Sakai (1997) recognised ten subfamilies; the Sarcinatricinae and Eudohrninae are additional to those recognised by Steinmann (1993). The adults are generally large, most often winged, and have a lobiform second tarsal segment. The most diverse subfamilies are the Opisthocosmiinae, Cosmiellinae, Forficulinae and Anechurinae. The family contains the ubiquitous tramp species, Forficula auricularia Linnaeus, to which a large literature is devoted; in Australia it is known from the cool temperate regions.

Nomenclature and spelling of family-group names of Dermaptera
Incorrect original spellings of family-group names and junior synonyms have had common usage in the dermapteran literature. The Anisolabididae are synonymous with Carcinophoridae and Psalidae, the Anisolabidoidea (= Carcinophoroidea), Spongiphoridae (= Labiidae, sic), and Spongiphoroidea (= Laboidea, sic). Senior synonyms are used in this work, even though the application of priority departs from the family-group names used by Popham (1965–1985) and Steinmann (1986–1993).

The family-group names based on genera which have the suffix, -labis have been routinely misspelled. In Cassis (1998), they are corrected in line with the recommendations of the International Code of Zoological Nomenclature (1985) and as listed in part by Sakai (1997) and Haas (1995). The suffix -labis is from the Greek, meaning forceps, whose stem is -idos and is feminine in gender. Therefore, the correct spelling of the relevant family-group names is as follows: Anisolabididae, Parisolabidinae, Brachylabidinae, Isolabidinae, Antisolabidinae, Platylabidinae and Anisolabidinae.

Unfortunately, no subsequent spelling correction has been made in the literature for the family-group Labidinae, which has been referred to by all other authors as the Labiinae (also Labiidae and Laboidea). The change may cause some instability in the literature but such corrections are required under Article 29b of ICZN 1985.

TAXONOMIC WORKS

Catalogues, checklist and bibliographies
A number of generic and species checklists of earwigs are available on a regional and global scale. These have been superseded by the world catalogue of Steinmann (1989a), although it has significant shortcomings.

There are also a number of checklists amongst the early literature. Scudder (1876a, 1876b, 1876c) and Kirby (1904) provided synonymic checklists that contained geographic descriptors and served as a framework for taxonomic work on earwigs. Burr's (1911) seminal work on dermapteran classification included a complete checklist of known genera and species, although few synonymical details were provided. Townes (1945) gave a checklist of genus-group names, which includes the type species and the method of designation.

Popham & Brindle (1966–1969) in a series of short papers, provided a checklist of genera and species under each family. These works include novel modifications to the family-group classification, commentary on previous works, new synonymies and keys to genera. They remain useful, despite the idiosyncracies of their suprageneric classification.

Sakai's (1970–1996) self-published series, the Dermapterorum Catalogus, serves as an unconventional checklist, but is barely decipherable and is not indexed in any reasonable fashion. It does, however, constitute a scientific publication for the purposes of nomenclature. It is chiefly a reorganisation of the literature, which has been photocopied and pasted together for each taxon. In more recent volumes, original colour photographs of species are provided. Most volumes have no index, pagination is not always sequential, the same taxa are treated in separate volumes without cross-referencing, and new synonymies are given without justification. Some volumes are little more than unexplained morphometric data. Nonetheless, his work brings much of the primary literature together, including many difficult references, and has some original work.

Sakai's (1982) checklist is his most useful work on the Dermaptera. It includes a review of previous classifications by Burr (1911), Popham (1965), Steinmann (1975) and Kevan (1977). Sakai's classification listed all known genera and species under each family, with general political keywords as descriptors of their distribution. This list recognises four superfamilies, 11 families, 49 subfamilies, 191 genera and 1798 species. The work is most useful for its bibliography which is more comprehensive than that of Steinmann (1989a).

Steinmann (1973, 1978, 1979, 1982a, 1982b, 1982c, 1983, 1984) published a series of checklists, which include illustrations of male genitalia, broad geographical distribution and synonymical listings. Steinmann's (1989a) world catalogue represents the most crucial document for any dermapterist. However, its value is diminished because of the selected bibliography, which is also a shortcoming of Steinmann's (1986, 1989b, 1990, 1993) monograph series. It should also be noted that the type information in Steinmann's world catalogue is often in conflict with his original works.

There are only a few regional catalogues of any significance. Srivastava's (1976) catalogue of the Oriental fauna is the most important of these works and has utility outside the region of interest. Reichardt (1968-1971) published a series of papers, cataloguing the Neotropical earwig fauna. Most other works of regional interest are taxonomic reviews (see below).

Taxonomic revisions
For bibliographic information for authors mentioned in this section, readers are referred also to Steinmann (1989a) and the Zoological Record 1989–1996.

Taxonomic work on earwigs is characterised by the efforts of very few, but nonetheless notable, dermapterists and the occasional forays of orthopterists. Most of the work has been carried out by Europeans, particularly by British entomologists throughout this century and the Hungarian dermapterist Henrik Steinmann.

Early works of significance include those of Scudder (1876a, 1876b) who provided diagnostic notes for many genera. Borelli (1888–1932) described numerous species from many parts of the world, particularly from Costa Rica and the Philippines. Borman's (1900) work was the first monograph of the Dermaptera and comprised two families (the Forficulidae and the Hemimeridae), 31 genera and 308 species. Dermapterology is indebted to the voluminous contribution of Captain Malcolm Burr, whose works spanned over a half century and 150 publications. His works were synthetic, modern and used a wide range of characters. They focused on the higher classification of earwigs and regional treatments or revisions of family groups.

Other British dermapterists include Walter Hincks, Edward Popham and Alan Brindle. Hincks (1935-1961) published over 50 papers on Dermaptera, his monograph of the Pygidicranidae (Hincks 1955, 1959) being the most notable. The majority of Popham's works (1959-1997) are concerned with morphology and higher classification. Brindle (1964-1988) published over 100 pages, regional in scope or taxonomic reviews of family-groups. His regional reviews of the earwigs of Madagascar, Surinam, the Afrotropical Region, the Solomon Islands, Caribbean, Sri Lanka, Micronesia and New Caledonia are some of the most important contemporary works on earwigs.

Bey-Bienko (1929-1970) provided the only significant taxonomic works on the earwigs of Russia and China. Srivastava (1969-1996) has published over 50 papers on the taxonomy of earwigs, primarily of the Oriental Region. Hebard (1917-1933) published extensively on the earwig fauna of the Neotropical Region and his work, together with the works of Brindle and Reichardt, offers some knowledge of the earwigs of this region.

Steinmann (1973-1993) has published over 50 papers on the taxonomy of earwigs. Many of his works are synthetic, culminating in his monumental monograph of the world's earwigs (Steinmann 1986, 1989b, 1990. 1993). This work provides keys and detailed descriptions of all taxa. Illustrations of the morphology of the species are restricted to the male and/or female forceps and the male genitalia where available. These works are essential for any dermapterist, although the prohibitive cost of each volume restricts their availability.

REVIEW OF THE AUSTRALIAN DERMAPTERAN FAUNA

Composition of the Australian dermapteran fauna
The described Australian dermapteran fauna comprises seven families, 18 subfamilies, 36 genera (Table 2) and 85 species. The Esphalmeninae (Pygidicranidae) and Opisthocosminae (Forficulidae) also occur in Australia (Victoria and Queensland respectively) although species have yet to be described. Despite Australia being a biodiversity hotspot for earwigs, only a third of the described subfamilies of earwigs have been recorded. Unlike a number of other insect orders, the degree of generic endemism among Dermaptera is very low in Australia, with only about 10% of the described genera restricted to the continent (cf. with Heteroptera with about 50% generic endemicity, see Cassis & Gross 1995). However, about half the described Australian species are endemic, and many new endemic species await description.

The Pygidicranidae are poorly represented in Australia, comprising less than 5% of the described fauna known from Australia, although undescribed species are known. Two of the genera, Austroblandex and Brindlensia, are endemic, and are placed within their own subfamilies. The Apachyidae are represented by three described species, and one undescribed species from the Northern Territory. The Labiduridae are also poorly represented, with only 4 of the 71 described species, known from Australia. The Anisolabididae in Australia are known from 10 described genera and 32 species, but many undescribed species are known to exist, particularly of genera such as Anisolabis and Gonolabis. The Spongiphoridae are represented in Australia by 10 described genera and 26 species and a number of undescribed species. The Forficulidae are poorly represented in Australia with very few species described or known in collections.

**Table 2.** Numbers of genera and species in the Demaptera recorded from Australia and the world. Endemic Australian taxa are given in parentheses.
	Australian		World
	Genera	Species	Genera	Species
Karschiellidae	0	0	2	12
Diplatyidae	0	0	5	161
Pygidicranidae	5 (2)	9 (5)	20	203
Apachyidae	1 (0)	3 (3)	2	15
Labiduridae	3 (0)	4 (2)	7	71
Anisolabididae	10 (2)	31 (19)	33	349
Spongiphoridae	10 (0)	26 (13)	39	512
Chelisochidae	4 (0)	8 (4)	16	96
Forficulidae	3 (0)	4 (2)	65	940
Total	36 (4)	85 (48)	189	1909

BIOLOGY
The biology of the Dermaptera is inadequately understood. Most information is related to species of economic importance, such as Forficula auricularia Linnaeus, or tramp species such as Labidura riparia. The most authoritative general accounts of dermapteran biology include those of Chopard (1938), Günther & Herter (1974), Albouy & Caussanal (1990) and Rentz & Kevan (1991).

The majority of earwig species are found in tropical and subtropical latitudes, although in Australia, many endemic species are found in cool temperate forests, with Anisolabididae commonly found in the litter. A number of tramp species, such as Anisolabis maritima, A. littorea and Labidura riparia are psammophilic. Earwigs are generally poorly represented in arid and semi-arid regions, although the minor pest, Nala lividipes, is known from semi-arid agroecosystems.

Earwigs are positively thigmotactic and are often found in secluded habitats, such as crevices and cracks, under bark, under stones, in litter and soil. They are also known to have moisture dependent microhabitat requirements and are negatively phototactic. Numerous species are synanthropic, commonly found in refuse, gardens, composts, and agricultural and industrial settings. Numerous species of Chelisochidae are known to occur in the leaf axils of Pandanus species in the Old World tropics.

Few generalisations can be made about the feeding habits of earwigs. Langston & Powell (1975) stated that in the tropics many species are obligate predators of small, soft-bodied insects, whereas in cooler regions earwigs appear to be mostly herbivorous, living on mostly dead or decaying vegetable matter. On limited evidence, there are indications that the Apachyidae and Chelisochidae are chiefly predaceous. Chelisoches morio is reported to feed on flies, leafhoppers, coccids, and holometabolous larvae. Labidura species are apparently predators of a wide range of arthropods; the Australian endemic Labidura truncata Kirby has been reported as predaceous on the codling moth Cydia pomonella (Linnaeus).

Omnivory is likely to be the most common feeding habit of earwigs. Anisolabis littorea and A. maritima are known to feed on crustaceans, such as isopods and amphipods, and on arthropod eggs. It is doubtful that earwigs are exclusively herbivorous, and known foodplants are mostly leafy vegetables. Nala lividipes has a wide range of foodplants that includes cotton, sunflower, sorghum, soybean and beetroot (Simpson 1989). Frugivory is also a suspected feeding habit, with Anisolabis dohrni (Kirby) known from the fruits of Artocarpus integer (Moraceae); Forficula auricularia is also recorded in association with fruit.

Acknowledgements

Dr David Rentz has provided significant support throughout the duration of the project and his efforts have greatly contributed to the successful completion of this work. Appreciation is extended to the Zoological Catalogue Series executive editor, Dr Keith Houston, who has maintained quality control throughout the project. The Trust of the Australian Museum, the Director, Dr Des Griffin, and Drs Doug Hoese and Mike Gray are acknowledged for institutional support of this work. The Head of the Division of Information Sciences, Ms Gwen Baker and her staff are especially thanked for their support in obtaining primary literature and information technology assistance. All but one of the illustrations used in the family introductions are from Rentz & Kevan (1991). They are reproduced with permission from CSIRO Entomology and the Melbourne University Press. Compilation of this section of the Catalogue was funded by the Australian Biological Resources Study.

Database Notes

The information on the Australian Faunal Directory site for the Dermaptera is derived from the Zoological Catalogue of Australia database compiled on the Platypus software program. It incorporates changes made to the work published on 2 September 1998 as (Cassis, G., 1998)

The classification used follows Haas (1995; Table 1). Within that, families, subfamilies, genera and species are arranged in alphabetic order, for self indexing purposes.

Family introductions
Family introductions indicate the possible sister-group relationships of the family and an indication of the number of genera and species worldwide. This is followed by a short description of the morphology of the family, which includes diagnostic features of the head, neck-structure, wings when present, and the male genitalia. The biological information is summarised and is often restricted to the described habits and habitats of well-known species. The suprageneric classification of the family is given, and includes, where applicable, a review of these taxa worldwide, regional distributions and any alternative taxonomic arrangements. The Australian fauna is summarised and gives the number of genera and species presently described, with some indication of the current status in work in preparation by the author on the systematics of the Australian earwigs. Details are provided about the most diverse Australian genera and any significant taxonomic works. Each introduction has a bibliography and a habitus illustration of a representative species of that family.

Synonymies
Complete synonymies for genera and species are given, including extralimital synonyms. All junior synonyms have been obtained from the primary literature or catalogues. Where different taxonomic arrangements exist, I have selected the most supported case and list the alternative views, including any putative junior synonym not accepted. Where there is multiple synonymy, each original authority is credited and often the authority of a monographic or revisionary work is also given. I have maintained strict adherence to ICZN (1985) as to what constitutes an available name.

New combinations
The authoroities for new combinations are given in the Reference field at the end of each species treatment. Alternative combinations can be traced through the currently accepted name and are not listed in the index.

References
Only lead taxonomic references for Australian Dermaptera are provided. Similarly there has been no attempt to provide a complete listing of biological and economic literature. Lead references have been provided, however, for pest species and will enable the user to access additional material. An attempt has been made to examine all the primary literature that contains original descriptions and new synonymies. Where the original literature was not examined, reference is made in the Catalogue to a secondary source.

Most entries in the Reference field have a brief explanation of the contents in parentheses after the reference and relevant page number(s).

Type specimens
All obtainable information on the primary types (holotypes, syntypes, lectotypes and neotypes) is provided, and includes reference to any subsequent designation. The depositories listed below are given for all type specimens and designated as seen or not seen (the latter marked by an asterisk).

Type localities
The type localities are given as in the original description, but where label data are different I have given priority to the information in the literature.

Distribution
This work contains information on the distribution, including extralimital distribution, of genera and species. Family-group distributions are generally provided in the family introductions.

Where a genus is not widely known in a region, the distribution of genus groups is given as major zoogeographical regions, which are sometimes qualified by political descriptors.

Species distributions are given firstly as standard drainage divisions (see format, Map: Australian Political and Geographic Regions), and then as States, Territories and Australian territorial islands. Islands such as the Torres Strait Islands (QLD) are included under their present state or territorial political administration.

Common names
Common names follow Naumann (1993) and original publications.

Ecological keywords and biological information
Ecological keywords are given in the Ecology field for each species. A list of keywords used in the database is given below. These include life history, feeding and habitat descriptors. Additional information is provided after the keywords in a telegraphic style or in the family Introductions. The authority for the ecological information is usually provided in the Reference field or in the original publication of the species.

Limital Area

Distribution data in the Directory is by political and geographic region descriptors and serves as a guide to the distribution of a taxon. For details of a taxon's distribution, the reader should consult the cited references (if any) at genus and species levels.

Australia is defined as including Lord Howe Is., Norfolk Is., Cocos (Keeling) Ils, Christmas Is., Ashmore and Cartier Ils, Macquarie Is., Australian Antarctic Territory, Heard and McDonald Ils, and the waters associated with these land areas of Australian political responsibility. Political areas include the adjacent waters.

Terrestrial geographical terms are based on the drainage systems of continental Australia, while marine terms are self explanatory except as follows: the boundary between the coastal and oceanic zones is the 200 m contour; the Arafura Sea extends from Cape York to 124 DEG E; and the boundary between the Tasman and Coral Seas is considered to be the latitude of Fraser Island, also regarded as the southern terminus of the Great Barrier Reef.

Distribution records, if any, outside of these areas are listed as extralimital. The distribution descriptors for each species are collated to genus level. Users are advised that extralimital distribution for some taxa may not be complete.

This work contains information on the distribution, including extralimital distribution , of genera and species. Family-group distributions are generally provided in the family introductions.

Where a genus is not widely known in a region, the distribution of genus groups is given as major zoogeographical regions, which are sometimes qualified by political descriptors.

Australian Biological Resources Study

Australian Faunal Directory